Geologica Acta, Vol.16 Nº 1, March 2018, 75-92 DOI: 10.1344/GeologicaActa2018.16.1.5 M. Bieńkowska-Wasiluk and M. Pałdyna, 2018 CC BY-SA
Taxonomic revision of the Oligocene percoid fish Oligoserranoides budensis (Heckel, 1856), from the Paratethys and paleobiogeographic comments
1 2 M. BIEŃKOWSKA-WASILUK AND M. PAŁDYNA
1Faculty of Geology, University of Warsaw Żwirki i Wigury 93, PL-02-089 Warszawa, Poland. E-mail: [email protected]
2Geoscience Friends Association ‘Phacops’ Grajewska 13/40, PL-03-766 Warszawa, Poland. E-mail: [email protected]
ABS TRACT Small perch-like fishes from the Oligocene of the Paratethys have been traditionally assigned toSerranus budensis (Heckel, 1856). A morphological revision of the holotype and specimens previously assigned to S. budensis from the Outer Carpathians, Poland, is provided herein. They are re-assigned to the species Oligoserranoides budensis (Heckel, 1856) — Percoidei incertae sedis. New specimens assigned to Ol. budensis from the Outer Carpathians, Poland, are introduced. Our results reveal that Ol. budensis is very similar to the species Caucasoserranoides morozkiensis, Carpathoserranoides brnoensis, Carpathoserranoides polonicus, and Oligoserranoides comparabilis, and further studies are necessary to revise the validity of those species. Ol. budensis shares many characters with genera Lutjanus and Ocyurus of the superfamily Lutjanoidea. Ol. budensis differs from Lutjanoidea in having a toothless palatine. The palaeobiogeography of Oligocene small perch-like fishes in the Paratethys is presented and discussed.
KEYWORDS Teleostei. Perciformes. Taxonomy. Morphology. Outer Carpathians.
INTRODUCTION However, Micklich (1998) indicated that this assignment was incorrect because of the absence of three spines on Small perch-like fishes, common in the Oligocene of the opercle of S. budensis, a diagnostic character of the Outer Carpathians, have been traditionally assigned the Serranidae (Johnson, 1983). Prokofiev (2009) and to the perciform species Serranus budensis (Heckel, Bannikov (2010) assigned S. budensis (referred in their 1856) (see Paucă, 1933; Jonet, 1958; Jerzmańska, 1968; papers to as Oligoserranoides budensis and Oliganodon Kotlarczyk et al., 2006). This species has also been budensis, respectively) to Percoidei incertae sedis due reported in the Oligocene of the Caucasus and the Upper to the lack of diagnostic characters of any fossil or extant Rhine Graben (Danil’chenko, 1960; Pharisat, 1991; percoid family, and noting the morphological differences Micklich, 1998; Pharisat and Micklich, 1998; Prokofiev, and similarities to some fossil and extant taxa. Prokofiev 2009; Bannikov, 2010). Recently, Prokofiev (2009) (2009) indicated the resemblance of Oligoserranoides selected S. budensis as the type species of his new genus to Symphysanodon (family Symphysanodontidae) and Oligoserranoides. While, Bannikov (2010) placed S. compared characters as the length of the snout, presence budensis in his new genus Oliganodon. The species S. of scales on the head, presence of the postmaxillary budensis was formerly assigned to the family Serranidae process, number of rays of the anal and the dorsal fin, by Danil’chenko (1960) and Jerzmańska (1968). number of vertebrae, predorsal formula and the number
75 M . B i e ń kowska-Wasiluk and M. Pa ł d y n a The Oligocene fish Oligoserranoides budensis from the Paratethys
of epuralia. Bannikov (2010) noted the resemblance of Bannikov (2006), and Jimtylerius temnopterus (Agassiz, Oliganodon to Symphysanodon and listed differences 1836) was classified as Percoidei incertae sedis by as the number of rays of the dorsal fin and the number Bannikov and Carnevale (2007). of abdominal vertebrae. In addition, Bannikov (2010) compared Oliganodon to the genera Pinjalo, The aims of this contribution are i) re-study the Parapristipomoides, Aphareus (family Lutjanidae) holotype (Fig. 1A) and fish material from the Outer and Hemilutjanus (family Serranidae, after Eschmeyer Carpathians of Poland previously referred to as Serranus et al., 2017). From the Oligocene of the Caucasus, budensis, ii) study the newly collected specimens of Danil’chenko (1960) described Serranus comparabilis, small perch-like fishes (Figs. 1B and 2), iii) redescribe latterly, Prokofiev (2009) assigned this species to his Oligoserranoides budensis, iv) compare Ol. budensis new genus Oligoserranoides, whereas Bannikov (2010) with some Palaeogene and living fishes, and v) study assigned it to his new genus Oliganodon. Under the the palaeobiogeographic distribution of Ol. budensis and principle of priority of the International Commission some others Oligocene percoid fishes in the Paratethys. on Zoological Nomenclature, ICZN, (1999), the name Oligoserranoides, introduced by Prokofiev (2009), is used herein for the species budensis and comparabilis. Material and methods
The large and diverse suborder Percoidei has been The holotype of Oligoserranoides budensis is housed inadequately defined and serves as a repository for in the Naturhistorishes Museum, Wien. The specimens representatives of the order Perciformes that cannot be previously referred to as Serranus budensis are from placed under any other suborder (e.g. Johnson, 1984). Prof. A. Jerzmańska’s collection (University of Wrocław) Percoids are diagnosed by a number of plesiomorphic from the Rogi and Winnica localities, and from collection characters. The fossil record of the Percoidei is scarce of University of Warsaw from Rudawka Rymanowska in comparison to their recent high diversity. The earliest locality. These localities are in the Outer Carpathians, percoid species based on articulated skeletons is from Poland. The new material (housed at the University the Cretaceous of India and is classified as Percoidei of Warsaw) comprises specimens collected by us and incertae sedis (Arratia et al., 2004). In contrast, the our collaborators in 1995–2015 in the Babice-Połanki, Cenozoic record includes many families defined from Dobra Góra, Futoma, Jamna Dolna, Jasienica Rosielna, articulated skeletons (see e.g. Blot, 1980; Patterson, Rudawka Rymanowska, and Winnica localities, in the 1993; Kotlarczyk et al., 2006; Bannikov, 2010; Carnevale Outer Carpathians, Poland. et al., 2014). The Cenozoic record includes many genera assigned to Percoidei incertae sedis due to the lack of Specimens from the Outer Carpathians (deposited in diagnostic characters of any fossil or extant family (e.g. the University of Wrocław and University of Warsaw) Bannikov, 2010; Carnevale et al., 2014), i.e. families are are from the Menilite Formation (Fm.) (Kotlarczyk, defined by many diagnostic characters, most of which et al., 2006) of the Silesian and Skole units, southern are of soft anatomy, thus, not possible to recognize in Poland. fossil specimens and it is not possible to assign such specimens to any family. In the Rogi (RO1 in Kotlarczyk et al., 2006) and Winnica (W in Kotlarczyk et al., 2006) localities (near Within the Oligocene record of the Percoidei Krosno town), the specimens were derived from the upper incertae sedis, a few species share many similarities part of the Menilite Fm. of the Silesian Unit, ichthyofaunal with Ol. budensis. For instance, Caucasoserranoides zone IPM2 (Jerzmańska, 1968; Kotlarczyk et al., 2006, morozkiensis Prokofiev, 2009, Oligoserranoides pg.: 66, Table 26). comparabilis (Danil’chenko, 1960) and Pirsagatia sytchevskayae Prokofiev, 2002. These species have In the Jasienica Rosielna locality (Wasiluk, 2013) near been recorded from the Caucasus (Prokofiev, 2002; Krosno town, the specimens were recovered in the upper 2009). Carpathoserranoides brnoensis Prokofiev, 2009 part of the Menilite Fm. of the Silesian Unit, ichthyofaunal and Carpathoserranoides polonicus Prokofiev, 2009, zone IPM2. known from the Outer Carpathians, are also similar to Ol. budensis. Some percoid fishes from the Eocene of Monte In the Rudawka Rymanowska locality in area of Bolca (Italy) share many similarities with Ol. budensis. Sanok town (Bieńkowska, 2004; RR in Kotlarczyk et al., Goujetia crassispina (Agassiz, 1839), Ottaviania mariae 2006; Bieńkowska-Wasiluk, 2010), in the Silesian Unit, (Sorbini, 1983), Ott. leptacanthus (Agassiz, 1839) and the specimens were from three exposures (RU 01, RU 02, Veranichthys ventralis (Agassiz, 1839) were assigned and RU 03, see Bieńkowska, 2004, Fig. 1) of the Tylawa to the Lutjanidae by Bannikov and Zorzin (2004) and Limestones and the adjacent shales, ichthyofaunal zone
Geologica Acta, 16(1), 75-92 (2018) 76 DOI: 10.1344/GeologicaActa2018.16.1.5 M . B i e ń kowska-Wasiluk and M. Pa ł d y n a The Oligocene fish Oligoserranoides budensis from the Paratethys
IPM2. Specimens were derived from the middle layers (1968), ichthyofaunal zone IPM2 (Kotlarczyk et al., 2006, of the Tylawa Limestones (see Ciurej and Haczewski, pg.: 66, table 26) above ichthyofaunal subzone IPM2-T. 2012; their samples of limestones RR 9/06, RR6A-C/06, SUB. (Trachinus event). Fig. 4). The Dobra Góra locality (DG in Kotlarczyk et al., The Jamna Dolna locality, near the city of Przemyśl 2006), near Sanok town and Babice-Połanki, (BP1 in
(Jerzmańska, 1968; Jerzmańska and Kotlarczyk, 1968; JAC Kotlarczyk et al., 2006) near Przemyśl city represents and JDG in Kotlarczyk et al., 2006) is in the Skole Unit. the Rudawka Tractionite Member of the Skole Unit, Specimens come from the Rudawka Tractionite Member, ichthyofaunal zone IPM2 (Kotlarczyk et al., 2006, pg.: lithological unit F-G (Fig. 2) of Jerzmańska and Kotlarczyk 66, table 26).
A
5mm
B
5mm
FIGURE 1. Oligoserranoides budensis holoypes. A) NHMW 1858/III/25. B) MWGUW ZI/57/014/b.
Geologica Acta, 16(1), 75-92 (2018) 77 DOI: 10.1344/GeologicaActa2018.16.1.5 M . B i e ń kowska-Wasiluk and M. Pa ł d y n a The Oligocene fish Oligoserranoides budensis from the Paratethys
5mm
FIGURE 2. Oligoserranoides budensis. Juvenile specimen, MWGUW ZI/57/051/b.
In the Futoma (Błażowa) locality near city of Rzeszów parhypural; pl, palatine; pmx, premaxilla; pop, preopercle; (Bąk et al., 2013), in the Skole Unit, the specimens ps, parasphenoid; pt, posttemporal; pts, pterosphenoid; were recovered from the Futoma Diatomite Member, pu, preural centrum; q, quadrate; ry, ray; ri, rib; r., right; ichthyofaunal zone IPM2. sc, scapula; scl, supracleithrum; SL, standard length; sn, supraneural bone (predorsal); soc, supraoccipital; sp, spine; Summing up, all studied specimens from the Outer sy, symplectic; uhy, urohyal; un, uroneural; v, vertebra; vo, Carpathians are Early Oligocene (Rupelian) in age. vomer; V, pelvic fin. Ichthyofaunal zone IPM2 is correlated with calcareous nannoplankton Biozone NP23 (see Kotlarczyk et al., 2006). Catalogue numbers of specimens are in Table 1. SYSTEMATIC PALAEONTOLOGY
Institutional abbreviations: MWGUW, Muzeum Order: Perciformes Bleeker, 1859 Geologiczne im. Stanisława Józefa Thugutta at the Suborder: Percoidei Bleeker, 1859 Faculty of Geology, University of Warsaw; NHMW, Family: incertae sedis Naturhistorishes Museum, Wien; ZPALWr., Prof. A. Jerzmańska’s collection, Department of Palaeozoology, GENUS Oligoserranoides Prokofiev, 2009 Faculty of Biological Sciences, University of Wrocław. Type of species. Smerdis budensis Heckel, 1856
For the anatomical terminology, we follow Johnson Diagnosis (emended). The genus is diagnosed by the (1981) and Potthoff et al. (1988). The anatomical term following unique combination of characters: maximum predorsal formula introduced by Ahlstrom et al. (1976, body depth in standard length 21-40%; supramaxilla pg.: 297) provides information on the position of the absent; palatine toothless; preopercle with serration; supraneural bones (predorsals) and the anterior dorsal opercle with two spines; 7 branchiostegal rays, ceratohyal pterygiophores in relation to the anterior neural spines (see without a beryciform foramen; posttemporal with Johnson, 1981). serrated posterior margin; 24 vertebrae (10 abdominal); three predorsals; predorsal formula 0/0/0+2/1+1/ or Anatomical abbreviations: aa, angulo-articular; A, /0+0/0+2/1+1/; 8 pleural ribs; pectoral fins long, reaching anal fin; br, branchiostegal ray; ch, anterior ceratohyal; cl, anterior part of anal fin and with 14-17 rays; dorsal fin cleithrum; co, coracoid; cr, crest; D, dorsal fin; d, dentary; continuous with 9 to 10 spines and 9 to 11 soft rays; three ecp, ectopterygoid; eh, epihyal (or posterior ceratohyal); ep, spines and 8 to 9 soft rays in anal fin; caudal fin forked epural; ept, endopterygoid; fr, frontal bone; hh, hypohyal; with 17 principal rays; three epurals; procurrent spur hp, hypural; hy, hyomandibula; in, interneural space; io, lacking; and ctenoid scales. infraorbital; iop, interopercle; l., left; leth, lateral ethmoid; mpt, metapterygoid; mx, maxilla; na, nasal; nsp, neural Remarks. The two anteriormost neural spines of the spine; op, opercle; P, pectoral fin; p, pterygiophore; ph, holotype are not preserved. Therefore, the predorsal
Geologica Acta, 16(1), 75-92 (2018) 78 DOI: 10.1344/GeologicaActa2018.16.1.5 M . B i e ń kowska-Wasiluk and M. Pa ł d y n a The Oligocene fish Oligoserranoides budensis from the Paratethys
bones are not clearly visible (Fig. 3B). It looks like the Diagnosis. As for genus. predorsal formula could be 0+0+0+2/1+1/, but the two neural spines should be near the supraneural bones. Material. MWGUW (46 individuals); ZPALWr. (33 The predorsal formula documented by Prokofiev individuals); for catalogue numbers of specimens see Table 1. (2009) is 0/0+0+2/ and /0+0/0+2/. The formula /0+0/0+2+1/1/ for Oligoserranoides comparabilis is Measurements. See Table 2. our interpretation from a figure provided by Bannikov (2010). The first predorsal formula (Fig. 4A) given in Description the present diagnosis is the most frequent in specimens from the Outer Carpathians. Both patterns of predorsal Skull formulae (0/0/0+2/1+1/ and /0+0/0+2/1+1/) occur in specimens from the same locality (e.g. Jamna Dolna, The skull bones of the holotype are poorly preserved, Jasienica Rosielna, Rogi, Rudawka Rymanowska). We with most of the bone margins unrecognizable have observed other patterns, such as the first given (Fig. 3A; B). The opercular bones and jaws are by Prokofiev and with variation of the position of moderately preserved. Most of the anatomical details two anterior neural spines on some specimens with were observed on specimens MWGUW ZI/57/015, signs of slight postmortem displacement of some head MWGUW ZI/57/028 (Fig. 3E; F), MWGUW ZI/57/036, bones or in which one of the anterior neural spines not MWGUW ZI/57/046 (Fig. 3C; D), ZPALWr. A/920, visible. ZPALWr. A/922. The head length is 33 to 42% SL. The frontal bones are smooth, without well-developed crests. Oligoserranoides budensis (Heckel, 1856) Figures 1-6 The limits of the skull roof bones and the position of the sensory canals are unclear. The parietal bones seem 1856. Smerdis budensis Heckel, pg.: 264, pl. XI, Fig. 16. to be about two times smaller than the frontals. The 1960. Serranus budensis Danil’chenko, pg.: 101, Fig. 21; supraoccipital crest is present but not well developed; pl. XXV, Fig. 4. specimen MWGUW ZI/57/046 has a serration preserved 1968. Serranus budensis Jerzmańska, pg.: 449, Fig. 19; on its posterior margin (Fig. 3C; D). Anteriorly, the pl. VI, Fig. 3. frontal sutures with the ethmoid region, but the 2009. Oligoserranoides budensis Prokofiev, pg.: 205, limits of the nasal bone and lateral ethmoid cannot Figs. 5-7 and 9. be determined. A narrow nasal bone is probably 2010. Oliganodon budensis Bannikov, pg.: 86, pl. VII, preserved in MWGUW ZI/57/046. Lateroanteriorly, Figs. 2 and 3. a large element, which we interpret as the lateral 2016. Oliganodon budensis Přikryl et al., pg.: 37, Fig. 4E. ethmoid, is present. The horizontal diameter of the orbit is 23 to 45% of the head length. The posterior, Holotype. NHMW 1858/III/25. anterior and ventral limits of the orbit are usually unclear. The parasphenoid is usually observed below Type locality and age. Blocksberg near Ofen, currently the middle region of the orbit, about the lower third Budapest (Schultz, 2013), Hungary; Oligocene. of the orbit.
TABLE 1. Studied specimens from the Outer Carpathians and its localities
Rogi ZPALWr. A/920–-923, A/925, A/1260, A/1262, A/1270–1272, A/1289–1290, A/1292, A/1294–1295, A/1308, A/1392, A/1437, A/1442–1443
Winnica ZPALWr. A/1467, A/1471, A/1474, A/1476–1477, A/1480, A/1483, A/1485, A/1487, A/1494, A/1507, A/1509–1510; MWGUW ZI/57/029
Jasienica Rosielna MWGUW ZI/57/016, ZI/57/035, ZI/5 7/039–41
Rudawka Rymanowska MWGUW ZI/57/018–20, ZI/57/022, ZI/57/028, ZI/57/033–34, ZI/57/042, ZI/57/047, ZI/57/053– 55, ZI/57/057–60, ZI/57/063–64, ZI/57/090, ZI/57/103, ZI/57/108–109, ZI/57/110/1
Jamna Dolna MWGUW ZI/57/014–15, ZI/57/036, ZI/57/04 6, ZI/57/048, ZI/57/050–51, ZI/57/104–107
Dobra Góra MWGUW ZI/57/065–67
Babice-Połanki MWGUW ZI/57/044
Futoma MWGUW ZI/57/023
Geologica Acta, 16(1), 75-92 (2018) 79 DOI: 10.1344/GeologicaActa2018.16.1.5 M . B i e ń kowska-Wasiluk and M. Pa ł d y n a The Oligocene fish Oligoserranoides budensis from the Paratethys
sn2 ?nsp2 ?nsp1 lines visible/distinct A B sn3 sn1 lines indistinct Dp1 pt fr lines reconstructed (on the specimen invisible) v3 ri scl hy pts P op ept r.sc? ps io pl r.pmx mpt ecp l.pmx cl l.sc? pop q mx r.co r.d l.?br l.co l.d
2mm br ch+hh l.cl r.?br
soc cr C D pt r.fr l.fr na? leth? vo? hy l.pmx io pl op cl ecp mpt d sy q l.mx pop
eh
2mm ch+hh
E F pt leth?
vo? r.pmx io l.pmx ept ecp l.op pl
r.q l.mx r.mx l.q pop l.aa pop l.d eh r.aa r.d iop ch hh uhy 5mm
FIGURE 3. Skull of Oligoserranoides budensis. A, B) holotype, NHMW 1858/III/25 photo (A) and camera lucida drawing (B). C, D) MWGUW ZI/57/046 photo (C), specimen is preserved as an imprint of bones (concave), photo made in a special technique to show a convex specimen, as if the bones were preserved, (D) interpretative drawing. E, F) MWGUW ZI/57/028/a photo (E) and drawing (F). Abbreviations: br, branchiostegal ray; ch, anterior ceratohyal; ch+hh, anterior ceratohyal plus hypohyal; cl, cleithrum; co, coracoid; d, dentary; Dp, dorsal pterygiophore; ecp, ectopterygoid; eh, epihyal; ept, endopterygoid; fr, frontal bone; hh, hypohyal; hy, hyomandibula; io, infraorbital; iop, interopercle; l.aa, left angulo-articular; l.br, left branchiostegal ray; l.cl, left cleithrum; l.co, left coracoid; l.d, left dentary; l.fr, left frontal; l.mx, left maxilla; l.pmx, left premaxilla; l.q, left quadrate; l.sc, left scapula; leth, lateral ethmoid; mpt, metapterygoid; mx, maxilla; na, nasal; nsp, neural spine; op, opercle; P, pectoral fin; pl, palatine; pop, preopercle; pt, posttemporal; pts, pterosphenoid; q, quadrate; r.aa, right angulo-articular; r.br, right branchiostegal ray; r.co, right coracoid; r.d, right dentary; r.fr, right frontal bone; r.pmx, right premaxilla; r.q, right quadrate; r.sc, right scapula; ri, rib; scl, supracleithrum; sn, supraneural bone (predorsal); soc cr, supraoccipital crest; sy, symplectic; uhy, urohyal; v, vertebra; vo, vomer.
Geologica Acta, 16(1), 75-92 (2018) 80 DOI: 10.1344/GeologicaActa2018.16.1.5 M . B i e ń kowska-Wasiluk and M. Pa ł d y n a The Oligocene fish Oligoserranoides budensis from the Paratethys
The infraorbital bones are usually no visible or not and with a straight postero-dorsal border. The preopercle preserved. Probably, they were thin and small. The has a serration on both the posterior and ventral margins lacrimal seems to be the largest infraorbital, but its shape (Fig. 4C; D). Small specimens have a serration and a few is unknown. Fragments of possible infraorbitals 2 or 3 are distinct spines where posterior and ventral margins meet preserved. (see Jerzmańska, 1968, fig. 19A; B). In larger specimens, serrations are slightly larger and more distantly placed at The hyomandibula lies vertically to the body axis. The the margin where the posterior and ventral margins meet. symplectic seems to be slim. The metapterygoid is only Specimen MWGUW ZI/57/046 (41.6 SL) has about 44 partially visible. The endopterygoid is large. The quadrate is preopercular spines in serration. Dorsal and ventral, i.e. triangular and meets anteriorly with the thin ectopterygoid, horizontal axes of the preopercle form an angle of slightly which does not project posteriorly to articulate with more than 90˚. the metapterygoid (see Johnson, 1981, figs. 21 and 22). Posterodorsal or posteroventral process of the quadrate is The hyoid arch is partially preserved. The epihyal not visible. The ectopterygoid articulates anteriorly with (or posterior ceratohyal) is triangular. The ceratohyal the long and thin palatine. Teeth or sockets for teeth on the (or anterior ceratohyal) does not present a beryciform endopterygoid, ectopterygoid and palatine have not been foramen. The number of branchiostegal rays in the observed. holotype is unclear, 6 or 7 rays are preserved. Specimens MWGUW ZI/57/015, MWGUW ZI/57/036, MWGUW The opercle has two small spines (Fig. 4B) and is ZI/57/046 have 7 branchiostegal rays. The number of much taller than wide. The limits of the subopercle are branchiostegal rays on anterior and posterior ceratohyals not visible. The interopercle is triangular, well developed, and epihyal is unclear. The anterior three rays are thinner
A nsp1 nsp2 B C D sn1 sn2 sn3
1 mm
1mm 2mm 2mm 2mm
E F G
E
1mm 1mm 1mm
FIGURE 4. Details of the skull and anterior vertebrae of Oligoserranoides budensis. A) anterior neural spines (nsp) associated with supraneural bones (sn), MWGUW ZI/57/065. B) opercle MWGUW ZI/57/014/b. C) preopercle MWGUW ZI/57/014/b. D) preopercle MWGUW ZI/57/029. E) premaxilla MWGUW ZI/57/033/a. F–G) jaws MWGUW ZI/57/033/a; area visible on E is indicated on F; G with outline of well-preserved teeth.
Geologica Acta, 16(1), 75-92 (2018) 81 DOI: 10.1344/GeologicaActa2018.16.1.5 M . B i e ń kowska-Wasiluk and M. Pa ł d y n a The Oligocene fish Oligoserranoides budensis from the Paratethys
A Dsp10 Dsp1-3 ?nsp2 Dry1-10 Dp1 sn2 sn3 ?nsp1
sn1 pt
in12 v3 scl
v10 ri op
r.sc? P cl l.sc? r.co Ar9 Ap1 l.co
Ary1
Asp1-3 l.cl 1mm
B C v10 v11
ri
Ap2
Ap1
2mm 2mm
FIGURE 5. Details of vertebral column, dorsal and anal fins of Oligoserranoides budensis. A) Camera lucida drawing of holotype, NHMW 1858/III/25, for interpretation of lines see Fig. 3B. B) Dorsal and anal fin MWGUW ZI/57/058 C) Anterior caudal region and anterior part of anal fin, ZPALWr. A/1494. Abbreviations: Ap, anal pterygiophore; Ary, anal ray; Asp, anal spine; cl, cleithrum; Dry, dorsal ray; Dsp, dorsal spine; in, interneural space; l.cl, left cleithrum; l.co, left coracoid; l.sc, left scapula; nsp, neural spine; op, opercle; P, pectoral fin; pt, posttemporal; r.co, right coracoid; r.sc, right scapula; ri, rib; scl, supracleithrum; sn, supraneural bone (predorsal); v, vertebra. than the posterior ones. The urohyal is triangular. Remains their reconstruction was impossible. Long gill rakers of ceratobranchials and epibranchials are preserved but were observed.
Geologica Acta, 16(1), 75-92 (2018) 82 DOI: 10.1344/GeologicaActa2018.16.1.5 M . B i e ń kowska-Wasiluk and M. Pa ł d y n a The Oligocene fish Oligoserranoides budensis from the Paratethys
The mouth is terminal, the snout length is usually smaller and postzygapophyses is unclear. We have observed than the horizontal diameter of the orbit. The premaxilla parapophyses on centra 6–10 (specimen MWGUW has well developed ascending and articular processes. The ZI/57/046); the posterior ones are the largest. The first caudal ascending process is sharp and narrow, slightly higher vertebra has a long hemal spine (Fig. 5A; C). The hemal than the articular process, which is rounded, not fused spines of preural centra 2–3 are thicker than the anterior ones along its anterior border with the ascending process. The and probably autogenous (specimen MWGUW ZI/57/028). postmaxillary process is broadly based, not finger-like. The neural spine of preural centrum 3 is longer and thicker The premaxilla usually bears small conical teeth, but some than the anterior ones. specimens have the teeth considerably enlarged anteriorly (Figs 3D, F; and 4E, G), other specimens seem to be Eight pairs of pleural ribs are articulated with vertebrae toothless. Because of a poor state of preservation, the teeth 3–10. On the holotype, the last pair is difficult to recognize. are not visible in the jaws of the holotype. The toothless maxilla is triangular, its depth increases posteriorly. The Some thin epipleurals are visible, but their number is supramaxilla is absent. unclear. Specimen MWGUW ZI/57/014, probably has 5 pairs of epipleurals associated with vertebrae 1–5. Intermuscular The lower jaw is moderately long and deep, and bears bones are absent in the caudal region. Three predorsal bones small teeth. The jaw articulation lies below the middle (supraneurals) are present. The first predorsal is usually of the orbit or slightly posteriorly (Fig. 3). The dentary anterior or posterior to the first neural spine. The second is a “V” shaped bone with an angle of “V” forming the predorsal is between the first and second neural spine. The symphyseal region. The angulo-articular is about the same third predorsal is posterior to the second neural spine. size as the dentary. The limits of the angulo-articular and the retroarticular are not visible. The pores of the mandibular Pectoral girdle and fin sensory canal are not well preserved. The post-temporal has a serrated posterior margin (see Vertebral column, ribs and intermuscular bones Fig. 3D; Prokofiev, 2009, fig. 7). Its intercalar process is slightly shorter than dorsal process for contacting the There are 24 vertebrae (including the preural centrum 1), epiotic. The supracleithrum is elongate and slim, its ventral 14 of which are caudal. In the holotype, the first vertebra portion overlaps the dorsal tip of the cleithrum (Figs. 3A is not preserved, but the tips of the first and second neural and 5A). The cleithrum is slightly S-shaped. It has a broad spines seem to be visible (Figs. 3B and 5A). The neural triangular-shaped shelf (posterior process) dorsally and a spines of abdominal centra 3–4 (Fig. 4A) are broader than broad shelf area ventrally. The ventral part ends sharply. the remaining spines. The neural spines of centra 9–11 are The postcleithrum is a well-developed, large and broad the tallest. Neural prezygapophyses are visible on centra bone with its ventral tip near the ventral margin of the body. 5–24 (specimen MWGUW ZI/57/046). The presence The boundary between postcleithra 1 and 2 is unclear. of neural postzygapophyses, haemal prezygapophyses The small scapula has a foramen (specimen MWGUW
ep1-3 un2 A B ep1-3 un2 C pu2 pu1 ry1 pu2 pu1 ?un2 hp5 hp4 ry8 hp3 ry9 hp3-5 hp2 hp3-5 ry8 hp1-2 hp1 hp1-2
ph ry1 ph 1mm 1mm 1mm
FIGURE 6. Details of caudal endoskeleton and fin of Oligoserranoides budensis. A) Camera lucida drawing of holotype, NHMW 1858/III/25. B) MWGUW ZI/57/016/a. C) MWGUW ZI/57/090/a. Abbreviations: ep, epural; hp, hypural; ph, parhypural; pu, preural centrum; ry, ray; un, uroneural.
Geologica Acta, 16(1), 75-92 (2018) 83 DOI: 10.1344/GeologicaActa2018.16.1.5 M . B i e ń kowska-Wasiluk and M. Pa ł d y n a The Oligocene fish Oligoserranoides budensis from the Paratethys
TABLE 2. Morphometric measurements of the studied specimens of Oligoserranoides budensis, and from Danil’chenko (1960), Jerzmańska (1968) and Bannikov (2010)
Holotype specimens specimens Danil’chenko Jerzmańska Bannikov NHMW MWGUW ZPALWr. (1960) (1968) (2010) 1858/III/25
SL (mm) 25.3 18.7-50.6 13-53 40-70 15.5-77 — *24 Maximum body depth in SL (%) 27.7 22-40 21-33 28-31 25-39.8 27-33 *33.3
Head length in SL (%) 33.6 31-43 34-38 33-35 — 33-36 *33.3 Horizontal diameter of orbit in head length (%) 23.5 23-45 29-35 — — — *25
* estimated if specimen was preserved with jaws closed
ZI/57/015). The coracoid is large, broad and deep, but it association and the third one in a serial association. The appears to be a thin bone. There are four small proximal second and third pterygiophore are placed together in the radials (specimen MWGUW ZI/57/015). fourth interneural space. Pterygiophores 4–10 are placed respectively in the succeeding interneural spaces. Two The pectoral fins attach relatively low on the flank, pterygiophores are placed in interneural spaces 12 and about the lower third of the body depth. They bear 15–20 14, and one pterygiophore is in interneural space 13. The rays each, which are long and reach the anterior-middle part remaining posterior pterygiophores are poorly preserved of the anal fin (hemal spine of fourth caudal vertebra). The and often they are displaced. The presence of three length of those fins cannot be established in the holotype segmental pterygiophores is unclear. and the number of rays is unknown. Anal fin Pelvic girdle and fin The anal fin bears three spines and 8 to 9 soft rays (9 The pelvic fins attach to the body just below the pectoral in the holotype, Fig. 4A). The first spine is the shortest, the fins or slightly posteriorly. They bear one spine and five third spine is the longest. rays each. Two elongate, triangular-shaped basipterygia (pelvic plates) are sutured to each other medially. The The first anal pterygiophore is enlarged and subpelvic and postpelvic processes are developed but their positioned more or less anteriorly to the hemal spine of shapes are unclear. The presence of accessory subpelvic the first caudal vertebra (Fig. 5A; C). Its triangular shape keel is unclear; most probably it is absent. is unclear in the holotype due to its poor preservation, because it is not in lateral view but is slightly rotated. The Dorsal fin dorsal tip of this pterygiophore often reaches the hemal arch of the last abdominal vertebra. Two supernumerary The dorsal fin is rather small, and bears 10 spines spines and one serially associated spine are present on the and 10 soft rays in the holotype (Fig. 5A) and in most first anal pterygiophore. The presence of three segmental of the studied specimens. There is a variability in the pterygiophores is unclear. number of rays in the dorsal fin bearing 9 to 11 soft rays and sometimes 9 spines. The fin is continuous, with the Caudal skeleton and fin spinous part not separated by a notch from the soft part. The posterior part of the fin (with soft rays) is usually The caudal fin consists of 17 principal rays (one slightly higher than the spinous one (Fig. 5B). The first unbranched, 7 branched in the lower lobe, 8 branched in dorsal pterygiophore or proximal radial 1 (Figs. 4A and the upper lobe and one unbranched). About 10 dorsal and 5A) is placed in the third interneural space (the first ventral procurrent rays are present. The caudal skeleton interneural space is between the skull and the first neural is represented by five hypurals, one parhypural and three spine) and bears two first spines in a supernumerary epurals (in the holotype the epurals are not preserved and
Geologica Acta, 16(1), 75-92 (2018) 84 DOI: 10.1344/GeologicaActa2018.16.1.5 M . B i e ń kowska-Wasiluk and M. Pa ł d y n a The Oligocene fish Oligoserranoides budensis from the Paratethys
the hypurals are poorly preserved; see Fig. 6). Details collection of Ol. budensis. from Rogi and Winnica, both of the first uroneural (the so-called stegural) and second small teeth and enlarged teeth were observed. Přikryl uroneural are not visible. Hypurals are distributed in two et al. (2016) noted slightly enlarged teeth around the groups (1+2; 3+4+5), which are separated by the hypural premaxillary symphysis (they ascribed their material to diastema. The neural spine of the second preural centrum Oliganodon budensis). is short. Neural spine of third, and hemal spines of second and third preural centra participate in the support of the Specimens ZPALWr. A/1266, A/1274, A/1350-1351, principal and procurrent caudal rays. The parhypural is as A/1359, A/1369, A/1375, A/1380, A/1386, A/1436, broad as the hemal spine of the second preural centrum. A/1440-1441, A/1458, A/1468-1470, A/1473, A/1475, Epurals are narrow and usually difficult to notice. A A/1482, A/1486, A/1488 assigned by Jerzmańska (1968) procurrent spur has not been observed in the holotype or to Serranus budensis probably belong to Ol. budensis in the additional material. according to our studies. However, they are too fragmentary or poorly preserved to justify their taxonomic assignment Scales to Ol. budensis. In our opinion, the above listed specimens should be classified as Percoidei incertae sedis. Scales are ctenoid, moderate in size, with ctenii at their posterior margins and a few radii (up to 12) in the anterior In this contribution, we have re-studied several fields. Ctenoid scales are observed on the trunk and cheek specimens published previously as Serranidae by (anteriorly to the preopercle, below the orbit, posteriorly Bieńkowska (2004, e.g. fig. 16) and Bieńkowska-Wasiluk to the region occupied internally by the ectopterygoid; (2010, e.g. fig. 28A, fig. 32 specimen a). According to our specimens MWGUW ZI/57/014, MWGUW ZI/57/028, present observations, some of those previously published MWGUW ZI/57/036, MWGUW ZI/57/046). Ctenoid specimens belong to Ol. budensis (specimens MWGUW scales seem to be present on the opercle (specimen ZI/57/018, ZI/57/028, ZI/57/033, ZI/57/034/1, ZI/57/042, MWGUW ZI/57/046). ZI/57/047, ZI/57/053-64, ZI/57/090, ZI/57/103, ZI/57/109, ZI/57/110/1), others should be assigned to Percoidei The lateral line is enclosed by scales. It is continuous, incertae sedis (specimens MWGUW ZI/57/017, ZI/57/021, concurrent with the dorsal profile, and descends to the ZI/57/032, ZI/57/037-38, ZI/57/043, ZI/57/045/1-2, level of the vertebral column near the twentieth vertebra ZI/57/056, ZI/57/061-62) due to the lack of diagnostic (unclear in the holotype). features, although they have an overall similarity to Ol. budensis, whereas the remaining specimens are in need of Pigmentation revision and are not discussed here.
Black and brown pigmentation appears on the dorsal part of the body (Fig. 5B; see Bieńkowska, 2004, fig. DISCUSSION 16; Prokofiev, 2009). Series of small melanophores are concentrated near the dorsal margin and disappear The small percoid fishes Caucasoserranoides gradually near the body midline. morozkiensis, Carpathoserranoides brnoensis, Carpathoserranoides polonicus, Oligoserranoides budensis, Ol. comparabilis and Pirsagatia sytchevskayae REMARKS from the Oligocene of the Paratethys have been described by Danil’chenko (1960), Prokofiev (2002, 2009) and Prokofiev (2009) illustrated the first pterygiophore of Bannikov (2010). the anal fin positioned posteriorly to the hemal spine of the first caudal vertebra. In the holotype ofOl. budensis and in The measured range in the body proportions of Ol. the studied Carpathian material, the first pterygiophore of budensis, i.e. maximum body depth in standard length, the anal fin is positioned anteriorly to the hemal spine of is wide and acquires 21–40%. This overlaps with the the first caudal vertebra (Fig. 5A; C). variability of Ol. comparabilis, i.e. 34–37% (Table 3). The range of this morphometric ratio for Ol. budensis according Only small teeth were mentioned in previous studies of to Bannikov (2010) is 27–33%. According to Danil’chenko Ol. budensis (see Danil’tshenko, 1960; Jerzmańska, 1968; (1960), Ol. comparabilis differs from Ol. budensis by Prokofiev, 2009; Bannikov, 2010). In the studied material, a longer head, larger maximum body depth and more many specimens are preserved as imprints of bones. In anterior origin of the dorsal fin. According to Bannikov such mode of preservation, the observation of teeth was (2010), in the dorsal fin Ol. comparabilis has 9 soft rays difficult. Sometimes the teeth seem to be small, but on the and Ol. budensis has 10 soft rays. Among the studied counterpart enlarged teeth are distinct. In the ZPALWr. specimens, there are individuals with 9 soft rays, there
Geologica Acta, 16(1), 75-92 (2018) 85 DOI: 10.1344/GeologicaActa2018.16.1.5 M . B i e ń kowska-Wasiluk and M. Pa ł d y n a The Oligocene fish Oligoserranoides budensis from the Paratethys
is no gap in the maximum body depth in standard length the Lutjanidae and to Ott. leptacanthus (Table 3). Both at 33–34% (Fig. 7). It is possible that Ol. budensis as we species are from the Eocene of Monte Bolca. Ott. mariae defined it herein includes alsoOl. comparabilis, but further resembles Ol. budensis in the body proportions, number studies of specimens of the latter species are necessary for of rays in dorsal and anal fins, and presence of ctenoidal resolving this issue. In the current interpretation, there is scales. It differs in the predorsal formula, presence of large no difference between the species Ol. budensis and Ol. conical teeth, fusion of hypurals 1–2 and 3–4, and presence comparabilis. of a broadly flanged last pleural rib. Ott. leptacanthus (see Bannikov, 2006) differs from Ol. budensis in the number of Oligoserranoides budensis shares many osteological abdominal vertebrae (but this character needs verification and meristic characters as well as measurements with and could be the result of postmortem processes), presence the Oligocene representatives of Carpathoserranoides of cycloid scales, and fusion of hypurals 1–2 and 3–4. Two and Caucasoserranoides. Carpathoserranoides species Goujetia crassispina and Veranichthys ventralis differs from Oligoserranoides in having 9 (versus 10) from the Eocene of Monte Bolca assigned to the Lutjanidae abdominal vertebrae. Caucasoserranoides differs from (Bannikov, 2006) significantly differ from Ol. budensis. Oligoserranoides in the predorsal formula (Table 3). The dorsal profile of the body being more convex than However, the variability in the predorsal formula may not ventral and the conical teeth distinguish both genera from be of taxonomic significance (see Ahlstromet al., 1976) as Ol. budensis. Both genera share the number of vertebrae it is assumed for Ol. budensis. Further studies are necessary and number of dorsal fin rays with Ol. budensis. Goujetia, in order to recognise, whether Carpathoserranoides, Jimtylerius, Ottaviania and Veranichthys were described Caucasoserranoides and Oligoserranoides are closely based on a very limited number of individuals that were related or even conspecific. much larger than Ol. budensis, which limits the possibility of comparison. Oligoserranoides budensis differs from Pirsagatia sytchevskayae in the predorsal formula, the number Oligoserranoides budensis is similar to family of branchiostegal rays, number of abdominal/caudal “Serranidae” Percoidei incertae sedis from the Palaeocene vertebrae (Table 3), and posteriorly inclined neural spines of Mexico (see Alvarado-Ortega et al., 2015) in the of the vertebrae (vs. neural spines in the middle part of the predorsal formula, number of spines and soft rays of the vertebral column, nearly vertically oriented). dorsal fin, number of vertebrae, number of spines in the anal fin, and presence of ctenoidal scales. It differs, however, in Prokofiev (2009) noticed the differences between Ol. the number of soft rays in the anal fin (8–9 vs. 7) and in the budensis and other perciforms such as Dapalis, Dapaloides absence of the supramaxilla. and Properca from the Oligocene of western Europe and Bilinia from the Oligocene-Miocene of the Czech Oligoserranoides budensis is similar in some characters Republic. Although Ol. budensis is defined in this paper in to Proserranus lundensis (Davis, 1890), a putative a few characters that are different than those suggested by perciform, from the Palaeocene (Danian) sediments of Prokofiev (2009), the differences presented by Prokofiev Limhamn, southern Sweden (Patterson, 1964). Ol. budensis (2009) between Ol. budensis and Dapalis, Dapaloides, shares with the latter species the number of vertebrae, Properca and Bilinia are significant. anal spines, dorsal spines and soft rays, and the presence of ctenoidal scales. The differences include a toothless With Jimtylerius temnopterus from the Eocene of Monte palatine and serration on the posttemporal posterior margin Bolca (Italy), Ol. budensis shares the predorsal formula, the in Ol. budensis. The lack of many anatomical data for number of dorsal and caudal fin rays, number of anal fin Proserranus does not allow for a more detailed comparison spines, number of vertebrae and body proportions (Table 3) of the species. (see Bannikov and Carnevale, 2007). Ol. budensis differs from J. temnopterus in the presence of ctenoidal scales, The presence of spines in the dorsal and anal fins, enlarged anterior teeth, and absence of a procurrent spur. In one spine and five rays (I, 5) in the pelvic fin, 17 both species, the hypurals are in two groups (1+2, 3+4+5) principal (I, 8, 7, I) rays in the caudal fin arranged separated by a diastema, the neural and haemal spines of on five hypurals and the parhypural, 7 branchiostegal the third preural vertebra are long, the neural spine of the rays and absence of the second ural centrum indicate second preural vertebra is reduced to a short crest, and there alignment of Ol. budensis with perciform fishes (see are three epurals. J. temnopterus was classified asPercoidei Johnson and Patterson, 1993). The number of rays incertae sedis by Bannikov and Carnevale (2007). in the pelvic and caudal fins is a primitive perciform character (Johnson, 1984). Ol. budensis presents the Oligoserranoides budensis is similar to Ottaviania characteristics of Percoidei (see Johnson, 1984). The mariae, assigned by Bannikov and Zorzin (2004) to presence of three predorsal bones, the predorsal formula
Geologica Acta, 16(1), 75-92 (2018) 86 DOI: 10.1344/GeologicaActa2018.16.1.5
M . B i e ń kowska-Wasiluk and M. Pa ł d y n a The Oligocene fish Oligoserranoides budensis from the Paratethys
TABLE 3. Comparison of some osteological characters and measurements of Oligoserranoides budensis and of some Palaeogene and extant fishes
Number of vertebrae Head Body Enlarged (number of Predorsal length in depth in Anal fin Dorsal fin anterior Age abdominal + formula standard standard teeth caudal length length vertebrae)
Oligoserranoides budensis 0/0/0+2/1+1/ Yes and this study 24 (10+14) III + 8-9 IX-X+9-11 and 31-43% 21-40% Oligocene no /0+0/0+2/1+1/
Oligoserranoides comparabilis 24 (10+14) III+8*1 X+9*1 /0+0/0+2+1/1/*2 No 32-37% 34-37% Oligocene acc. to Bannik ov (201 0)
Caucasoserranoides 0/0+0+2/1+1/1/ morozkiensis and 36.4- 26.7- 24 (10+14) III+8-9 X+10 Yes Oligocene acc. to Prokofi ev (200 9) 0/0+0+2/1/1/*3 39.4% 27.3%
Carpathoserranoides brnoensis 0+0+0/2/1 and ca. 34% acc. to Prokofi ev (200 9) 24(9+15) III+9 X+10 /0+0+0/2/1/ No (2.9 times 42% Oligocene in SL)
Carpathoserranoides polonicus ca. 37% 4 5 acc. to Prokofi ev (200 9) 24(9+15)* III +ca. 8 X+10? /0+0+0/2+1/1* No (2.7 times - Oligocene in SL)
Pirsagatia sytchevskayae Oligocene acc. to Prokofiev (200 2) 25(11+14) III + 9 IX + 10 0/0+0/2/1 No 36.4% 33.3% - Lower Miocene
Jimtylerius temnopterus acc. to Bannikov and Carneva le 24(10+14) III+7 X+9 /0+0/0+2/1+1/ No 31% 31% Eocene (2007)
Ottaviania mariae 24(10+14) III+8 X+11 /0+0+0/2/1+1/ Yes 34% 30% Eocene acc. to Bannikov and Zorzin (200 4)
Ottaviania leptacanthus 24(11+13) III+8 X+11 - - 33% 30% Eocene acc. to Bannikov (2006)
Symphysanodon 0/0/0+2+1/1/1/ acc. to Anderson (1970); 25(10+15) III+7-8 IX + 10 and Yes 27-36% 20-36% extant Johnson (1984) 0/0/0+2/1+1/1/
Lutjanus cyanopterus acc. to McEachran and Fechhelm 24(10+14) III+7-8 X+14 0/0/0+2/1+1/ Yes 36-37% 29-32% extant (2005); Fahay (2007)
Ocyurus chrysurus acc. to Alle n(198 5); McEachran 24(10+14) III+8-9 IX–XI +12–14 0/0/0+2/1+1/ Yes ca. 29% 30-34% extant and Fechhelm (2005); Fahay (2007)
*1 According to Danil’chenko (1960) for Serranus comparabilis. *2 From fig. 45 of Bannikov (2010). *3 The first predorsal formula is from the text and the second from fig. 1 of Prokofiev (2009). *4 In the text (Prokofiev, 2009) 9+15=25 vertebrae are cited, whereas 24 vertebrae are mentioned for the genus. *5 From fig. 15 of Prokofiev (2009).
0/0/0+2/, 24 vertebrae, three anal spines, the presence (Serranidae). Ol. budensis shares with Lutjanoidea many of five hypurals, three epurals, one ural centrum, and a characters, such as, ctenoid scales, scales on cheek, a low neural crest on the second preural centrum of Ol. complete lateral line, the number of dorsal and anal fin budensis are primitive features of the Percoidei (see spines and soft rays, a single dorsal fin, the number of rays Johnson, 1984). The predorsal formula /0+0/0+2/1+1/ is in caudal, pectoral and pelvic fins, a serrated preopercle, a derived character and is present in the Acropomatidae, an opercle with two spines, a serrate posterior margin of which, however, differ in the presence of two dorsal fins the posttemporal, a premaxilla with larger conical teeth, (see Johnson, 1984; Fahay, 2007). the supramaxillary bone being absent, a ectopterygoid without teeth, the general anatomy of the suspensorium, Bannikov (2010) indicated the similarity between and the number of branchiostegals, vertebrae and pleural Ol. budensis and some genera of the family Lutjanidae ribs, three predorsals and the predorsal formula, the (Pinjalo, Parapristipomoides, Aphareus) and Hemilutjanus procurrent spur being absent, three epurals, and the number
Geologica Acta, 16(1), 75-92 (2018) 87 DOI: 10.1344/GeologicaActa2018.16.1.5 M . B i e ń kowska-Wasiluk and M. Pa ł d y n a The Oligocene fish Oligoserranoides budensis from the Paratethys
2525 L. carponotatus (Richardson, 1842), L. lutjanus Bloch, 1790 and L. madras (Valenciennes, 1831) in a similar number of 2020 Ol. comparabilis dorsal fin spines and soft rays, number of anal fin soft rays
1515 and body depth in SL (see Allen, 1985). The difference is the b vs. Ol. budensis bbbb forked ( emarginated) caudal fin of . It is similar 1010 in body proportions to the juveniles of several species of the
Maximum body depth body Maximum Lutjanidae, e.g. Etelis oculatus (Valenciennes, 1828), L. 55 cyanopterus (Cuvier, 1828) and Ocyurus chrysurus (Bloch, 00 1791) (see Clarke et al., 1997; Fahay, 2007; Victor et al., 0 1010 2020 3030 4040 5050 6060 7070 bbbSL 2009). Ol. budensis differs from the juveniles of Oc. chrysurus of subfamily Lutjaninae in a forked (vs. emarginated) caudal FIGURE 7. Measurements variability of Oligoserranoides budensis, fin (adults of Oc. chrysurus do have a forked caudal fin). The maximum body depth and standard length. The line indicates 34% maximum body depth in standard length for Ol. comparabilis. difference between Oc. chrysurus and Ol. budensis is the number of soft rays in the dorsal fin (Table 3). The assumed variability of Ol. budensis in the number of dorsal and anal of hypurals (see Johnson, 1981). Ol. budensis differs from fin rays is comparable to such variability in extant species the superfamily Lutjanoidea in having a toothless palatine. of the Lutjanidae (see Allen, 1985). The pigmentation of Ol. Maybe Ol. budensis is a member of the superfamily budensis resembles L. cyanopterus (see Clarke et al., 1997) of Lutjanoidea, but further detailed studies of the osteology comparable size but is clearly different from L. campechanus of the species and the representatives of the superfamily (Poey, 1860), L. griseus (Linnaeus, 1758) and L. synagris (both fossil and extant) are necessary. (Linnaeus, 1758) (see Fahay, 2007).
Further research is necessary especially for the Oligoserranoides budensis differs from the Sparoidea observation of characters typical for the Lutjanoidea such in having seven (vs. 6) branchiostegal rays and a different as presence of trisegmental posterior 1-7 dorsal and anal predorsal formula. Ol. budensis differs from the Haemuloidea pterygiophores, the shape of the lacrimal, presence of the by having 24 (vs. 26-27) vertebrae, the absence of the subocular shelf, presence of well-developed metapterygoid procurrent spur, and a different predorsal formula. lamina, details of the suspensorium, inserting of the branchiostegal rays on the ceratohyal and epihyal, details Oligoserranoides budensis resembles the extant of the branchial skeleton, details of the neurocranium, and Symphysanodon (family Symphysanodontidae; number and articularion of epipleurals. Prokofiev, 2009; Bannikov 2010) in body proportions, meristic characters, the presence of a single dorsal fin, Oligoserranoides budensis shares with the Lutjanidae ctenoidal scales (Table 3), and suspensorium anatomy. the number of dorsal and anal fin spines and soft rays, Symphysanodon differs in 15 caudal vertebrae, the moderately protrusible premaxillaries, an ascending predorsal formula and two (out of three) strongly reduced process not a separate ossification, a broadbased and not epurals (Prokofiev, 2009). fingerlike postmaxillary process, and the presence of canines in the upper jaw. GEOGRAPHIC AND STRATIGRAPHIC DISTRIBUTION Oligoserranoides budensis differs from the subfamily Etelinae (family Lutjanidae), which include Aphareus, in The holotype of Ol. budensis is from Budapest, the predorsal formula. Hungary, in the Inner Carpathians, and the species was mentioned by Weiler (1933) in the Oligocene of Hungary Oligoserranoides budensis differs from the Apsilinae (a more precise age was not indicated). Specimens of Ol. in the number of dorsal fin spines and soft rays and the budensis occur in the Outer Carpathians, Caucasus, and in number of anal fin soft rays. the Upper Rhine Graben (Fig. 8).
Oligoserranoides budensis differs from the subfamily In the Outer Carpathians of Poland Ol. budensis is Lutjaninae (family Lutjanidae) which include Pinjalo, noted frequently from the Lower Oligocene, Menilite Fm., the subfamily Paradichthyinae (family Lutjandae) and the ichthyofaunal zone IPM2, and occurs in the Upper Oligocene family Caesionidae in the absence of a posterior projection in the same formation in ichthyofaunal zone IPM6 (Kotlarczyk on the ectopterygoid. et al., 2006). Serranus sp., which probably represents Oligoserranoides was noted in the entire succession of the Oligoserranoides budensis resembles Lutjanus Menilite Fm. correlated with the Oligocene-Lower Miocene, bengalensis (Bloch, 1790), L. boutton (Lacepède, 1802), in ichthyofaunal zones IPM1-7 (Kotlarczyk et al., 2006). The
Geologica Acta, 16(1), 75-92 (2018) 88 DOI: 10.1344/GeologicaActa2018.16.1.5 M . B i e ń kowska-Wasiluk and M. Pa ł d y n a The Oligocene fish Oligoserranoides budensis from the Paratethys
material studied herein is from ichthyofaunal zone IPM2, Pirsagatia sytchevskayae is derived from the Upper which is correlated with the middle Rupelian, calcareous Oligocene or the Lower Miocene of the Caucasus nannoplankton Biozone NP23 (see Kotlarczyk et al., 2006). (Prokofiev, 2002).
Oligoserranoides budensis is also known from the Difficulties in the correlation of the Oligocene Oligocene of the Outer Carpathians from the Czech deposits from the Upper Rhine Graben, Outer and Inner Republic (Prokofiev, 2009), Romania (Paucă, 1933; Jonet, Carpathians, and the Caucasus (see Popov et al., 2002; 1958, under the name Serranus budensis) and from Ukraine Prokofiev, 2009; Bannikov, 2010) do not allow for a (Prokofiev, 2009; Bannikov, 2010). more precise analysis of the geographic and stratigraphic distribution of Ol. budensis. In the Upper Rhine Graben, Ol. budensis was described from the Lower Oligocene, from the Froidefontaine locality in France, in marls with Meletta (Marnes à Mélettes) (Pharisat, PALAEOBIOGEOGRAPHY 1991), as well as from coeval deposits (Meletta-Schichten) of the Lower Oligocene from the Frauenweiler locality in The fossil record of Ol. budensis indicates its wide Germany (Micklich, 1998; Pharisat and Micklich, 1998). distribution in the Paratethys (see Fig. 9) in the Lower Oligocene and in the lower Upper Oligocene strata. In the Caucasus, Ol. budensis is noted from the upper In contrast, Ol. comparabilis, known only from the Lower Oligocene and lower Upper Oligocene, at the Belaya Lower Oligocene of the eastern Paratethys, seems River, Morozkina Balka Horizon, and at the Gumista River to be the Caucasian endemic species that was not a (Bannikov, 2010). competition of Ol. budensis. Ol. comparabilis occurs earlier than Ol. budensis in the Early Oligocene in the Ol. comparabilis is known from the Lower Oligocene eastern Paratethys. In such case, the migration of Ol. of the Pshekhian Horizon, Caucasus (Danil’chenko 1960; budensis from the Carpathian region to the eastern Bannikov 2010). Paratethys seems to be possible. Caucasoserranoides morozkiensis and Pirsagatia sytchevskayae seem Caucasoserranoides morozkiensis was described from to be endemic species from the eastern Paratethys, the upper Lower Oligocene and lower Upper Oligocene of whereas Carpathoserranoides brnoensis and the Morozkina Balka Horizon, Caucasus (Prokofiev, 2009). Carpathoserranoides polonicus were endemic from the Carpathian Region of the Paratethys. Presence of Carpathoserranoides brnoensis and Carpathoserranoides species known only locally is in concordance with polonicus were described from the Oligocene of the Outer limited connections of the Carpathian Region and Carpathians (Prokofiev, 2009). the eastern Paratethys (Fig. 9). In contrast, the wide
Oligoserranoides budensis Oligoserranoides comparabilis Moravia Poland Ukraine Carpathoserranoides brnoensis Germany Carpathoserranoides polonicus France Hungary Caucasoserranoides morozkiensis Caucasus Pirsagatia sytchevskayae Romania
FIGURE 8. Geographic occurence of the Oligocene small percoid fishes.
Geologica Acta, 16(1), 75-92 (2018) 89 DOI: 10.1344/GeologicaActa2018.16.1.5 M . B i e ń kowska-Wasiluk and M. Pa ł d y n a The Oligocene fish Oligoserranoides budensis from the Paratethys
Rafał Nawrot, Radosław Wasiluk, Robert Czerwiński and Piotr Gryz for helping in the fieldwork; Grzegorz Komendowski for photographic documentation of some
Polish Ukrainian specimens, Anna Żylińska for correcting the English writing. Carpathians Carpathians Frauenweiler Moravia CARPATHIAN We are very grateful to Gloria Arratia and an anonymous Froidefontaine BASIN reviewer for reviewing the manuscript and improving the Budapest PARATETHYS Romanian Caucasus style. This work was supported by funds of the Faculty of Carpathians Geology, University of Warsaw, DSM grant 105518 to M. Bieńkowska-Wasiluk.
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Manuscript received September 2016; revision accepted September 2017; published Online January 2018.
Geologica Acta, 16(1), 75-92 (2018) 92 DOI: 10.1344/GeologicaActa2018.16.1.5